Crustaceana 46 (2) 1984, E. J. Brill, Leiden

ON THE TAXONOMIC STATUS OF TIGRINA EYDOUX & SOULEYET, 1842 (, BRACHYURA)

BY B. GALIL and CH. LEWINSOHN t Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Israel

INTRODUCTION Trapezia tigrina, first described by Eydoux & Souleyet, 1842, from Hawaii, has since been reported, mistakenly, as a synonym of a number of allied species. This misidentification stemmed from an original description lacking in some critical taxonomic features and misplacement of the type. Thus, re- description was deemed necessary to permit accurate identification of this tax- on in future studies. The various synonymies are discussed. Trapezia tigrina Eydoux & Souleyet, 1842 (fig. 1) Cancer rufopunctatus - Ruppell, 1830: 27, 28 (mentioned under T. coerulea). Trapezia tigrina Eydoux & Souleyet, 1842: 232, pi. 2 fig. 4; Ward, 1939: 13, figs. 15, 16. Trapezia maculata - Dana, 1852: 256; 1855: pi. 15 fig. 4; Streets, 1877: 103; De Man, 1888: 319, pi. 13 figs. 2, 2a; Henderson, 1893: 366; Alcock, 1898: 221 (p.p.); Nobili, 1906: 293; Stimpson, 1907: 73; Baiss, 1924: 13; Ramadan, 1936: 35; Monod, 1938: 142; Edmondson, 1946: 301, fig. 180f. - Jacquinot & Lucas, 1853: 41 (p.p.); Heller, 1861a: 13; Heller, 1861b: 350; Hilgendorf, 1869: 75, pi. 2 fig. 3b (p.p.); A. Milne-Edwards, 1873: 258 (p.p.); Kossmann, 1877: 42 (p.p.); De Man, 1880: 176; De Man, 1881: 94; Miers, 1884: 536 (p.p.); Klunzinger, 1913: 309, pi. 7 fig. 13. Trapezia ferruginea var. rufopunctata - Paulson, 1875: 48, pi. 7 figs. 3-3a. Trapezia ferruginea maculata - Ortmann, 1897: 206 (p.p.); Lenz, 1912: 4. Trapezia ferruginea var. maculata - Borradaile, 1900: 590; Borradaile, 1902: 265. Grapsillus maculatus - Rathbun, 1906: 865. Trapezia cymodoce maculata - Rathbun, 1911: 235; Rathbun, 1930: 558, pi. 228 figs. 3, 4; Ed- monson, 1962: 300, fig. 31b. Trapezia cymodoce var. maculata - Laurie, 1915: 462. Trapezia danae Ward, 1939: 13, figs. 17, 18. Trapezia danai - Knudsen, 1967: 51; Garth, 1974: 402. Trapezia aff. danai - Serene, 1969: 136, figs. 14a, 17, 20 (p.p.); Garth, 1971: 198. Trapezia wardi Serene, 1969: 140; Serene, 1971: 914; Preston, 1973: 470; Edwards & Emberton, 1980: 235; Coles, 1980: 283. Trapezia aff. wardi - Garth, 1971: 198. Trapezia flavopunctata - Chen & Lan, 1978: 281, fig. 12. ?Trapezia rufopunctata - Cano, 1889: 211; Wedenissow, 1894: 413; Edmondson, 1923: 20. ?Trapezia rufopunctata var. maculata - Ortmann, 1893: 484; Guinot, 1962: 240. ? Trapezia ferruginea maculata - Lenz, 1901: 466; Lenz, 1910: 552. ?Trapezia maculata - Nobili 1901: 16; Nobili, 1905: 10; Lenz, 1905: 351; Laurie, 1906: 410; Stebbing, 1910: 304; Gravier, 1920: 470; Pesta, 1928: 72. TRAPEZIA TIGRINA 167 ?Trapezia cymodoce maculata - Rathbun, 1907: 52 (p.p.). ?Trapezia ferruginea forme maculata - Bouvier, 1915: 272. ?Trapezia ferruginea subsp. maculata - Michel, 1965: 30. not Trapezia acutifrons - A. Milne Edwards, 1867: 281. not Trapezia rufopunctata - A. Milne Edwards, 1873: 258. not Trapezia tigrina - Serene, 1969: 133. Material examined Red Sea: Elath, Gulf of Elath ( = Aqaba), 10 October 1952; la, cl. 10.2 mm; identified as T. wardi, "Type" by Serene (TAU NS 62); 10 October 1952; la, 19 cl. 10.1, 11.6 mm; identified as T. wardi, "Cotype" by Serene (TAU); 21 May 1958; lO*, cl. 13.4 mm (Senck. M. 4102). Nabeq, Gulf of Elath, Sinai; 17 October, 1979; la, cl. 7.1 mm (TAU). Gulf of Elath, Sinai; 24 June 1976; 2 m., on Stylophora; 1 a 1 9, cl. 10.8, 10.8 mm (TAU); 1980; on live ; 5(70* 999, cl. 10.9, 9.0, 7.5, 6.9, 6.7, 12.4, 10.7, 10.6, 10.5, 10.2, 10.0, 10.0, 9.8, 9.2 mm (TAU). Senafirld., entrance to Gulf of Elath, 8January 1949; coll. "Manihine"; la 399 ljuv-> cl. 12.2, 15.4, 13.4, 14.1, 9.2 mm (BM). Na'ama, N. of Sharm el Sheikh, Sinai; 15 October 1979; lo*, cl. 12.2 mm (TAU); 6 November 1981; 1 m deep from living , la 19, cl. 10.5, 11.3 mm (TAU). Sharm Moiya, N. of Sharm el Sheikh, Sinai; 3 February 1949; fauna assoc. with coral; coll. "Manihine"; la 1 9, cl. 11.3, 12.4 mm (BM 1959: 1:17:56-63). Et Tur ( = Tor), Gulf of Suez, Sinai; January-July 1826; leg. E. Ruppell; la 19, cl. 12.3, 12.1 mm (Senck. M. 4038). Daedalus Shoal, 1874; coll. Lt. Col. Playfair; 2aa 299 2 juv., cl. 12.8, 12.2, 14.1, 11.3, 7.6, 7.5 mm (BM 1874: 89). Shab Baraja, Sudan; 19 October 1977; on reef corals; 1 9 , cl. 14,8 mm (Senck. M.). Assab, Ethiopia, Red Sea; 15 February 1966; half dead coral, 50 cm; 2aa 2 99, cl. 12.2, 12.1, 13.8, 13.5 mm (RMNH). Red Sea; leg. R. Kossman; 5aa 999, cl. 11-6, 10.9, 10.8, 10.6, 9.9, 13.5, 13.3, 13.0, 12.7, 12.3, 12.0, 11.7, 11.3, 11.2 mm (Senck. M. 9548). Indian Ocean: Sapper Bay, Aden; October 1966; coll. K. England; la, cl. 9.6 mm (BM 1974: 99). Aden: 1885; la 4 juv., cl. 13.3, 6.6, 6.1, 5.2, 4.9 mm (BM 1885: 14). Gesira, 20 km S. of Mogadiscio, Somalia; August 1976; leg. M. Vannini; la 1 9, cl. 10.9, 11.9 mm (MF); October 1979; leg. M. Vannini; la 1 9, cl. 8.2, 11.6 mm (MF); la 1 9, cl. 12.1, 11.4 mm (MF); 19 ljuv.cl. 7.3, 4.3 mm (MF); 19 cl. 7.6 mm (MF); 4 juv., cl. 7.0,6.7, 3.7; 2.9 mm (MF); 5 juv., cl. 6.8, 6.1, 3.7, 3.5, 2.9 mm (MF). Etoile Island, Amirante Isl.; April 1882; 13 fms. coral; H.M.S. "Alert"; 2aa 1 9, cl. 11.7, 10.9, 12.2 mm (BM 1882: 24). Imma Is., Male Atoll, Maldives; 21 March 1964; ex ; 1 9 1 juv., cl. 7.9, 5.2 mm; (identified as aff. wardi Serene) (AH). Dunidu Is., Male Atoll, Maldives; 19 April 1964; ex. Pocillopora woodjonesi; la 1 9 , cl. 15, 18 mm (AH). Pacific Ocean: Phillipines; 15 January 1908; U.S.F.C. Str. "Albatross"; la 1 juv., cl. 11.2, 5.4 mm (USNM). Ifaluk Atoll, Caroline Isl.; 1953; la, cl. 10 mm (USNM 200652). Kapingamarangi Atoll, Caroline Isl.; 12 August 1954; 1 9 , cl. 12 mm (USNM 230088). Eniwetok Atoll, Marshall Isl.; 18 August 1966; coll. J. W. Knudsen; la 19, cl. 11.7, 13.4 mm; identified as T. aff. danai Serene, 1969 (AH). Rigili Is., Eniwetok Atoll, Marshall Isl.; 2 August 1966; coll. J. W. Knudsen; 1 a , cl. 7.4 mm (AH). Japtan Is., Eniwetok Atoll, Marshall Isl.; 4 August 1966; coll. J. W. Knudsen; 2 99, cl. 5.7, 4.4 mm (AH). 170 B. GALIL & CH. LEWINSOHN Honolulu, Hawaii; 1834; 18 fms; H.M.S. "Challenger". 19, cl. 15.7 mm (BM 1834: 31). Waikiki, Oahu, Hawaii; 1923; 1 9, cl. 16.6 mm (BPBM S 1568). Kauai, Hawaii; 1902; U.S.F.C. Sta. "Albatross"; 1(7 19 1 juv., cl. 13.5, 15.8, 6.1 mm (USNM 29486). Pt. Allen, Kauai, Hawaii; 12 October 1959; 8crcr 2 99, cl. 14.8, 14.9, 12.5, 13.1, 12.1, 12.0, 9.2, 7.8, 10.8 mm (BPBM S 6610). Anaa Atoll, Tuamotu Arch.; 27 October 1967; 60-70 ft.; National Geographic Soc.; lcr, cl. 11 mm (USNM 291828). Tikehae Lagoon, Tuamotu Arch.; 11 April 1957; lcr 1 9, cl. 9.0, 9.0 mm (USNM 213821). Society Isl.; coll. A. Garrett, id. W. Faxon; lcr 29 9, cl. 10.9, 11.4, 7.7 mm (USNM 158983). South coast of Pitcairn Is., S. Pacific; 20 October 1967; 22 fms. coral; National Geographic Marquesas Expedition; la 1 9, cl. 12.8, 14.8 mm (USNM 291828); 20 September 1967; 22-25 fms.; coral; National Geographic Marquesas Expedition; 1 O*, cl. 7.8 mm (USNM 291828), largest male, cl. 15 mm; female, cl. 18 mm; smallest juvenile, cl. 2.9 mm. The material is deposited in the Allen Hancock Museum in the University of Southern California (AH); the British Museum (Natural History) (BM); Ber- nice P. Bishop Museum, Honolulu (BPBM); Rijksmuseum van Natuurlijke Historie, Leiden (RMNH): Museo Zoologico of the University of Florence (MF); Senckenberg Museum (Senck. M.); the Zoological Museum of Tel Aviv University (TAU) and the U.S. National Museum (USNM). The abbreviation cl. stands for carapace length. Morphological description. — Carapace smooth, slightly convex, without definite regions. Antero-lateral margins slightly curved inwards in the direc- tion of outer orbital spine, but inflated in the middle so that the margins are parallel along the proximal half. In younger specimens (cl.<8 mm) the carapace is quite as broad at the outer angles of the orbits as between the lateral teeth or even slightly divergent, but in larger specimens, especially gravid females, it is slightly contracted in front. Posterolateral margins convergent backwards from a distinct lateral epibranchial tooth which in juveniles always projects as a sharp spine, some adults retain that character. Front, four-lobed, projects beyond the orbits, over the antennules and bases of the antennae which are excluded from the orbits. The frontal submedian pair of lobes are triangular, separated from each other by a shallow V-shaped incision and from the outer pair of lobes by a wider, uneven sinus; both lobes of this outer pair are about twice as wide basally as each submedian lobe, and directed obliquely outwards, their edge sometimes finely denticulate. Apices of outer pair of fron- tal lobes not extending as far forward as the submedians. The internal angles of the orbits are rounded and less prominent than the front. The orbits, too shallow to conceal the eyes, are cut out of the antero-lateral angles of the carapace. The inferior inner orbital tooth is conical, incurved and rather pro- minent. The postorbital tooth is acute, often incurved. The antennae have the peduncular articles small, fitted in between the outer side of the slender anten- nular groove and the inner side of the inferior orbital angle. The third max- illipeds are subrectangular, with the exognath not quite extending to the distal angle of the endognath. The ischium of the endognath with the inner margin TRAPEZIA TIGRINA 169

Fig. 1, Trapezia tigrina Eydoux & Souleyet. A, female (cl. 11.3 mm) from Na'ama, Sinai, Red Sea; B, male (cl. 12.2 mm) from Assab, Ethiopia, Red Sea; C, male (cl. 11.5 mm) from Aden, south coast of Arabian Peninsula; D, male (cl. 12.1 mm) from 20 km S of Mogadiscio, Somalia; E, female (cl. 12.5 mm) from Kauai, Hawaii. rounded. Inner lip of merus rounded, inner distal angle truncate and outer distal margin rounded. Chelipeds smooth, glabrous and considerably developed. Chelipeds subequal in females and young males, noticeably une- qual in adult males. The meri project beyond the lateral margins of the carapace; the distal margin is provided with one triangular tooth. Anterior margins armed with 5-7 small teeth, simple or serrulate, which increase in size 170 B. GALIL & CH. LEWINSOHN distally. The shape and number of the meral teeth is subject to variation, often the right and left arms of a specimen are strongly unequal. Minute teeth, observed only under magnification, decorate the lower margin of the palm. In- ner angle of carpus produced, covered with squamiform granules. A tooth-like pointed prominence decorates the antero-internal angle of the carpus. Fingers incurved, distally acute. Ambulatory legs slender, with segments slightly com- pressed. Dactyli sparsely pubescent and rather shorter than propodi. The distal end of the dactyl is curved and composed, on the upper surface, of several rounded ridges, blunted distally, thus giving the tip an appearance of a denticled rasp. Posterior margin in ambulatory dactyli with five parallel transverse rows of short, blunt spines and a cluster of stout setae at distal end. The shape and number of rows is of importance systematically. Color in alcohol. — Conspicuous round, reddish spots on a creamy yellow or rosy background cover carapace, abdomen and pereiopods. The configura- tion of the spots varies among juveniles and adults. Remarks. — The type of Trapezia tigrina Eydoux & Souleyet, 1842, is deposited in the Museum National d'Histoire Naturelle in Paris. Despite a passage of 140 years, the dry specimen is in good state of preservation, even though all traces of color have disappeared. Its box, No. 2950, contained the legend: " T. flavomaculata Eydoux & Souleyet. Trapezie taches jaunes, type uni- que" and inside a handwritten note by J. Forest, 1960, "je doute que ce soit le type de T. jlavopunctata car il ne correspond pas aux dimensions du type." Another box, No. 2968, which contains a specimen of T. Jlavopunctata bore the legend: "T. tigrina type", and a note by D. Guinot: "il y a une erreur dans les types de Trapezia Jlavopunctata et T. tigrina. Le type No. 2968 de tigrina est en fait celui de Jlavipunctata de la fig. PI. 2 fig. 3 d'Eydoux et Souleyet espece bien con- servee outre sa couleur par la bord inferieur de la paume du chelipede avec 'des tubercules' dit Eydoux et Souleyet, fortement denticule comme la figure et le type 2968. Le type No. 2950 de flavomaculata (erreur pour Jlavopunctata) est en fait celui de tigrina de la PI. 2 fig. 4, espece conservee outre sa couleur par le bord inferieur de la paume des chelipeds en ' 'crete (lisse) trachante"." And in- deed, checking measurements given by Eydoux & Souleyet and their excellent figures, I have no doubt that the contents of those boxes were exchanged either by the original authors or not long afterwards: A. Milne Edwards (1867), while describing T. acutijrons, writes: "Cette espece se rapproche beaucoup par sa taille de la Trapezia tigrina Eydoux & Souleyet". T. acutijrons A. Milne Edwards is accepted as a synonym to T. rujopunctata which indeed bears a close resemblance to T. Jlavopunctata. that probably by then occupied the box of T. tigrina. Serene (1969) too, clearly erred when he described T. tigrina as having "the frontal lobes well marked but comparatively less salient and less acute than rujopunctata and acutijrons; the lower margin of palm of cheliped strongly granular". Figure 4 in his manuscript given as "tigrina (Type, Paris Museum)" is obviously that of Jlavopunctata. TRAPEZIA TIGRINA 171 The unfortunate exchange together with a typographical error in Eydoux & Souleyet (1842) which placed the figure of T. tigrina at number 2 instead of 4 were probably the source of ambiguity concerning this species. Dana (1852) was first to include T. tigrina*) into synonymy with T. maculata (Macleay), not his fault really, as Macleay's description (1838) consisted of just three lines. Thus, many subsequent authors followed Dana in the use of the name maculata: Streets (1877), Ortmann (1897), Alcock (1898) and Lenz (1901), wrote of T. maculata (Macleay) Dana. Unfortunately, Macleay's type is lost now. Mrs. Helen Tranter of the Australian Museum, Sydney wrote us: "The position is that the Macleay Museum no longer appears to have the type of this species and there is no record of there ever being any paratypes, ...The locality of the type must now be regarded as unknown". However, it seems that T. maculata (Macleay) is close to T. rufopunctata. Ward (1939) examined Macleay's type and then went on to establish a new species T. danae for T. maculata as described by Dana, 1852, (not T. maculata (Macleay, 1838)). His description lacks in details, but photographs were taken. Barnard (1950: 278) writes "from the photograph of McLeay's specimen sent me by Mr. Ward, it appears that Krauss' surmise that this was rufopunc- tatus was correct. The photograph shows traces of large spots on the cheliped, which has a serrulate lower border". Alas, this photograph too was lost. D. Griffin & P. Stabury (1970) examined the type and identified it as T. rufopunc- tata. J. S. Garth (1974), who checked this type too, agrees with this identifica- tion. Ward (1939) established T. danae n. sp. for T. maculata as described by Dana. T. danae was separated from T. maculata (Macleay) by an elongated carapace, less developed frontal teeth and smaller spots on carapace and chela. Though Ward overlooked the character of the lower border of the palm which is serrulate in T. maculata Macleay, it is evident from the included photographs (figs. 17, 18) that in this specimen the palm is entire. The American Museum of Natural History possesses Ward's (1939) specimens of T. tigrina and T. danae type specimens from Samoa and kindly let the senior author (B.G.) examine them. T. danae Ward corresponds in all morphological aspects to T. tigrina Eydoux & Souleyet. Ward placed his specimens of T. tigrina near T. rufopunctata (Herbst) and then listed differences concerning the shape of frontal teeth, lack of serrulation on manus and proportion of carapace which clearly described his specimens as true T. tigrina Eydoux & Souleyet. Ward's figs. 15 and 16 are of a typical juvenile T. tigrina. Serene (1969) described T. aff. danai from material in Allan Hancock Museum collected by J. Garth. He based his identification on the denticulation of the anterior border of the merus, a notoriously unstable character in

*) Dana's specimens are not extant but his description and drawings are lucid and we con- fidently assign them to T. tigrina. 170 B.GALI L& CH. LEWINSOHN Trapezia. As comparative material, Serene chose specimens from Mauritius (his figs. 14a, 21) which are decorated with more than 120 spots on the carapace, while the type of T. danae Ward has 50 spots. Upon close examina- tion by us the specimens from Mauritius were assigned to T. richtersi, a new species described by us not long ago. Thus, we can reject Serene's observations on the number of spots decorating the carapace of T. danae. Serene (1970) on Vietnamese material established T. wardi n. sp. for T. tigrina as described by Ward. He separated it from T. danae by its having a wider carapace, an acute external orbital angle and epibranchial teeth, the merus of the cheliped is armed with 6 or 7 sharp teeth terminating in outcurved spines - all features actually characterizing young specimens of T. tigrina. Serene wrote that T. tigrina, as described by Ward, is different morphologically from the type of T. tigrina Eydoux & Souleyet (1842) which he had examined in the Paris Museum, but as mentioned above, he was misled by the interchange of Eydoux & Souleyet's types. In the Allan Hancock Museum of the University of Southern California, we found the specimens decribed by Serene (1969) as T. wardi. We checked too, material from Elath (Gulf of Aqaba, Red Sea) identified by Serene as T. wardi "type" and "cotype" but not published as such; these specimens are il- lustrated in figs. 17-19 in Serene's (1969) paper. Serene (1969) saw mostly very small specimens, none exceeding 8 mm, in his own words: "The specimens of the present collection are too small, mainly the male, to be identified with some certitude but they are closer to wardi than to danai \ In our opinion, Serene mistook the juvenile features of T. tigrina and thus established a new species. After careful examination of the literature and a considerable amount of material from many parts of the Indo-West Pacific, we are of the opinion that the material and the synonymies mentioned here belong to a single valid species - T. tigrina Eydoux & Souleyet, 1842. Acknowledgements. — Thanks to Mr. Z. Wolodarsky for help assembling a collection from the Gulf of Elath. We wish to express our gratitude for en- trusting us with material to Dr. D. M. Devaney, Bishop Museum; Dr. J. S. Garth, Allan Hancock Museum; Dr. R. W. Ingle and Mr. P. Clark, British Museum; Dr. R. B. Manning, U.S.N.M.; Dr. M. Tiirkay, Senckenberg Museum and Dr. M. Vannini, University of Florence. Photographs were taken by Ms. L. Maman. This work is part of a Ph.D. thesis by B. Galil. ZU S A M M EN F ASS U N G Trapezia tigrina Eydoux & Souleyet, 1842, wurde nach Material aus Hawaii beschrieben. Die Art geriet in Vergessenheit und spateres Material von ihr wurde mit anderen Arten der Gattung identifiziert, hauptsachic.h mit T. rufopunctata (Herbst, 1799) und T. maculata (Macleay, 1838). Serene (1970) beschrieb die Art als neu unter dem Namen T. wardi. Zu den genannten Irrtii- mern fuhrten hauptsachlich zwei Grunde: Die ungeniigende Originalbeschreibung der Art und die Verlegung des Typusexemplares von T. tigrina in die Schachtel des Typusexemplares von T. flavopunctata Eydoux & Souleyet und umgekehrt. Die Art wird hier neu beschrieben und die verschiedenen Synonyme diskutiert. TRAPEZIA TIGRINA 173

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Received for publication 28 July 1982.